Duellman, William E., Marion, Angela B. & Hedges, Blair, 2016, Phylogenetics, classification, and biogeography of the treefrogs (Amphibia: Anura: Arboranae), Zootaxa 4104 (1), pp. 1-109: 52

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Hylidae  : Cophomantinae

Shortly after the origin of the clade herein referred to as cophomantines, an ancestral stock inhabited the earlyuplifted western part of the continent (the proto Andes). This clade representing current Colomascirtus  , Hyloscirtus  , and Myersiohyla  diverged (crown node) in the mid-Eocene 47.5 (38.0–57.0) Mya. These frogs inhabited streams, and their tadpoles developed large suctorial mouths with many rows of labial teeth as they dispersed through the ever-rising Andes. An early divergence (the crown node) in northern South America restricted Myersiella  to the Guiana Highlands, whereas in the Andes in the early Oligocene Colomascirtus  differentiated from Hyloscirtus  33.3 (29.3–37.3) Mya. Coloma et al. (2012) estimated the minimum divergence time of these events as 61.2 and 40.9 Mya, respectively. The latter inhabited low to moderate elevations, whereas Colomascirtus  came to inhabit streams at higher elevations.

The other major clade within Cophomantinae subsequently diverged into two lineages in the late Eocene, 36.8 (32.8–40.8) Mya. One clade consisted solely of Bokermannohyla  , which inhabited the mountains of southeastern Brazil and developed stream-adapted tadpoles. In the other clade, a lowland lineage, Hypsiboas  , differentiated from a highland group, Aplastodiscus  with stream-adapted tadpoles, in the late Eocene, 34.2 (30.9–37.5) Mya.

The earliest divergence within Hypsiboas  occurred in the mid-Oligocene 31.9 (28.4–35.5) Mya when a large group of species ( H. benitezi  H. sibleszi  ) occupied northeastern South America including the Guiana Highlands. Differentiation of most clades of Hypsiboas  occurred in the Miocene. For example, the long-legged tree frogs of the Hypsiboas albopunctatus  Group differentiated about 17.4 (14.0– 20.8) Mya in the Amazon Basin and dispersed into coastal Brazil. Likewise, the ancestral large, arboreal frogs in Hypsiboas albomarginatus  Group diverged about 19.2 (15.4–22.9) Mya. Other species groups are more recent. For example, the clade of Andean species in the Hypsiboas balzani  Group differentiated in the Pliocene about 5.0 (13.8 – 6.1) Mya. Similarly, the Pliocene was the time of divergence of the ancestral stock of the Hypsiboas pulchellus  Group, 5.7 (4.8–6.5) Mya that came to inhabit subtropical highlands from Bolivia to southern Brazil.